Minnesota Invasive Terrestrial Plants & Pests Center New Species Evaluation

Lupinus polyphyllus Lindl. (Large-leaved lupine)

Evaluated: A.C. Morey; Reviewed: R.C. Venette (1/15/20)

OVERVIEW:

Common names: bigleaf lupin(e), Washington lupine, marsh lupine, garden lupine, meadow lupine Synonyms: see Beuthin (2012) and CABI (2018) for varieties, cultivars, and synonyms

Lupinus polyphyllus is a perennial legume (Fabaceae) native to the western U.S and Canada. It was introduced to the eastern U.S. (and Canada) as a garden plant, but is also used for soil improvement and animal forage. It has also been introduced to Europe, New Zealand, and Australia and is considered a problem outside of cultivation in many regions. Lupinus polyphyllus is the dominant parent of the popular cultivated hybrid known as Russel Lupin or L. x regalis. Some sources note that garden escapees are likely L. x regalis misidentified as L. polyphyllus due to the greater relative prevalence of the hybrid in commercial trade.

MAJOR KNOWLEDGE GAPS ASSOCIATED WITH ASSESSMENT:

 Distribution within Minnesota  Economic impact (to yield/marketability, from quarantine/control)  Extent of future invasion  Impact to threatened/endangered species

1 | L. polyphyllus

ARRIVAL Proximity to Minnesota: VERY HIGH

RANKING Very High Pest is known to occur in Minnesota Pest occurs in Wisconsin, Iowa, South Dakota, North Dakota, Manitoba or High Ontario Medium Pest occurs in North America Low Pest is not known to occur in North America

Lupinus polyphyllus is currently reported outside of cultivation in at least seven (7) counties in Minnesota (Cook, Lake, St. Louis, Itasca, Carlton, Pine, and Becker) (EDDMapS 2019; UMN-Bell 2019), with the earliest herbarium record occurring in 1970 in Itasca Co. (UMN-Bell 2019). Other sources suggest additional counties have populations (Benton and Aitkin) (Chayka and Dziuk 2017; Kartesz 2015).

It has otherwise been documented in regions of Wisconsin, Michigan, Alaska, and Maine (EDDMapS 2019), and likely in additional northeastern states (USDA-NRCS 2019; Kartesz 2015). It is also introduced in Ontario and other eastern Canadian provinces (USDA-NRCS 2019; Canadensys 2019).

Existence of Pathways: HIGH

RANKING High Pathways for arrival of the pest in Minnesota are known to occur Pathways for the arrival of the pest in Minnesota are conceivable, but not Medium known to occur Low Pathways for arrival of the pest in Minnesota are difficult to conceive

Lupinus polyphyllus is documented within Minnesota (see Proximity to Minnesota), and no regulation currently exists to limit its arrival or spread to the state.

It is a species historically and presently part of the horticultural trade used as a garden ornamental, for erosion control, soil improvement, as an intercropping species, as forage for wildlife and livestock, and in fire-protection belts (CABI 2018; Beuthin 2012; Fremstad 2010). Seeds and plants of L. polyphyllus or its hybrids (Russel lupine) or cultivars (‘Gallery Pink’) are widely available for purchase online (e.g., amazon.com, etsy.com, buyrareseeds.com, bulkwildflowers.com, brecks.com), including through at least one local retailer (plants.gertens.com).

Entry into Minnesota could also occur through the natural dispersal of viable seeds or vegetative fragments (see Innate Dispersal Capacity); Lupinus polyphyllus is found in nearby regions in WI, MI, and Ontario (see Proximity to Minnesota).

2 | L. polyphyllus

Innate Dispersal Capacity: LOW

RANKING Maximum recorded dispersal >500 km per year (or moves in low level Very High jets/ upper atmosphere) High Maximum recorded dispersal 500-250 km per year Moderate Maximum recorded dispersal 100-250 km per year Maximum recorded dispersal 1-100 km per year (wind dispersal; flowing Moderately Low water) Maximum recorded dispersal <1 km per year (movement through soil; Low splash dispersal)

Lupinus spp. seeds are comparatively heavy and do not have specialized structures for wind dispersal or adhesion to animals (OGTR 2013). They are dispersed mainly through ballistic dispersal, shooting away from parent plants by a few meters (OGTR 2013; Ramula 2014). There is suggestion that L. polyphyllus seeds are dispersed via zoochory by insects and/or rodents (Rask-Jensen 2018), though evidence is lacking. Lupinus polyphyllus can also spread locally through creeping rhizomes (Fremstad 2010; Ramula 2014).

Besides human-mediated spread and potential zoochory, long distance dispersal of L. polyphyllus seeds is said to occur via waterways (OGTR 2013; CABI 2018), but documented evidence of this could not be found.

ESTABLISHMENT AND PERSISTENCE

Suitability of Minnesota Climate: HIGH

RANKING High >40% of Minnesota is predicted to be suitable Medium >20 to 40% of Minnesota is predicted to be suitable Low >0 to 20% of Minnesota is predicted to be suitable Negligible No part of Minnesota is suitable

Documentation of the climatic tolerance of L. polyphyllus is limited. It prefers “cool, subalpine climates” (Beuthin 2012) and perennial Lupinus x hybrida, the most popular of which are L. polyphyllus-based “Russel Hybrids”, are listed as USDA Zone 4-8 plants (MBG 2019). Records in Glacier Co., MT confirm its tolerance to at least Zone 4, though most of its native range in the western U.S. occurs in warmer Zones (USDA- NRCS 2019; USDA-ARS 2012).

Specific sites of L. polyphyllus in Becker, Itasca, and Pine Co., MN confirm occurrence of the species in USDA Zone 3b (EDDMapS 2019; USDA-ARS 2012). As a Zone 3b species, >40% of Minnesota is climatically suitable.

3 | L. polyphyllus

Presence of Hosts: HIGH

RANKING High >10% of Minnesota with suitable hosts (or habitat for weeds) Medium >1 to 10% of Minnesota with suitable hosts (or habitat for weeds) Low >0 to 1% of Minnesota with suitable hosts (or habitat for weeds) Negligible 0% of Minnesota with suitable hosts (or habitat for weeds)

In its native western U.S. range, L. polyphyllus occurs primarily in meadows, roadsides, banks, and prefers seasonally wet habitats (CABI 2018; Fremstad 2010; Beuthin 2012).

Habitats of L. polyphyllus in its invaded range are often associated with disturbance, but can vary, including open ruderal areas, such as road verges, wastelands and meadows, forest understories, river banks, and wetlands (CABI 2018; Ramula 2014; Fremstad 2010). The species tolerates acidic, nutrient-poor soils and prefers moderately dry, sandy soils (CABI 2018; Fremstad 2010). Descriptions of sites were it has been document in Minnesota include dry and moist roadsides, disturbed fields, open grassy or rocky areas, and mixed wooded areas (UMN-Bell 2019; EDDMapS 2019).

Considering only non-interstate roadways, there are ~123,394 miles in Minnesota (Williams 2005). Assuming ‘roadside’ habitat could occur within 100ft of the road edge on either side, that equates to 4,674 square miles, or ~5.7% of total state land. In 2016, 1,058,955 acres were classified as CRP land (MN-DNR 2019), which is about 2% of total state land. There are ~4,420 miles of railroads in Minnesota (MN-DOT 2018; MRRA 2020). Assuming ‘railway’ habitat could occur within 100ft of the rail edge on either side, that is about 167 square miles, or ~0.2% of total state land. There are 4,455,584 acres of non-native prairie grasslands in the Prairie Region of Minnesota (MN-DNR 2018), which is about 8.7% of state land. Of the estimated 7,719,200 acres of riparian land in Minnesota, 34.5% is forested (2,663,124 acres) (MN-DNR 2001). This accounts for about 5.2% of Minnesota’s terrestrial land.

Only portions of these selected habitats would need to be suitable to account for >10% of Minnesota.

Hybridization/Host Shift: HIGH

RANKING High Species reported to hybridize or has undergone a documented host shift Medium Species in the same genus have been reported to hybridize/shift hosts Low Hybridization/Host shifts have not been reported for this genus or species

Species within Lupinus are known to hybridize, both naturally and through artificial breeding (Stace 2010; Clements et al. 2008; Kurlovich, Stoddard, and Earnshaw 2008).

4 | L. polyphyllus

Popular ornamental cultivars known as Russel Lupine or Russel Hybrids (= L. x regalis or L. polyphyllus ssp. polyphyllus) are based on hybridization between L. arboreus and L. polyphyllus (Fremstad 2010; Stace 2010; Beuthin 2012). This hybrid has been found spontaneously occurring in Britain where both parents occur, potentially backcrossing to L. arboreus (Stace 2010; BRC 2019). In Europe, spontaneous hybrids of L. polyphyllus with naturalized L. regalis and L. nootkatensis have been reported (KEW 2019; Fremstad 2010). Hybrids between L. polyphyllus and L. nootkatensis (=L. x pseudopolyphyllus) have also been reported in Alaska (CABI 2018).

Some sources state that L. polyphyllus will hybridize with L. perennis, a native eastern U.S. species and single host plant to the endangered Karner Blue butterfly (Byleckie 2000; WI-DNR 2019). The resulting hybrid is purportedly unsuitable for larval development. However, other sources are less certain of this phenomenon (Fremstad 2010) and empirical evidence of hybridization or unpalatability of such hybrid to the endangered butterfly could not be found.

SPREAD

Existence of Pathways: HIGH

RANKING High Pathways for arrival of the pest in Minnesota are known to occur Pathways for the arrival of the pest in Minnesota are conceivable, but not Medium known to occur Low Pathways for arrival of the pest in Minnesota are difficult to conceive

Intentional and unintentional human-mediated spread is considered to be a major long-distance pathway for L. polyphyllus movement into new areas (Brobäck 2015; CABI 2018; OGTR 2013), and naturalized populations of the species have been noted within Minnesota for more than 5 years (see Proximity to Minnesota).

It is a species historically and presently part of the horticultural trade as a garden ornamental, erosion control, soil improvement, as an intercropping species, as forage for wildlife and livestock, and in fire-protection belts (CABI 2018; Beuthin 2012; Fremstad 2010; Kurlovich, Stoddard, and Earnshaw 2008). Seeds and plants of L. polyphyllus or its hybrids (Russel lupine) or cultivars (‘Gallery Pink’) are widely available for purchase online (e.g., amazon.com, etsy.com, buyrareseeds.com, bulkwildflowers.com, brecks.com), including through at least one local retailer (plants.gertens.com).

5 | L. polyphyllus

Dispersal Capacity-Reproductive Potential: HIGH

RANKING High Annual reproductive potential (r) of pest is >500 descendants per year Medium Annual reproductive potential (r) of pest is 100 to 500 descendants per year Low Annual reproductive potential (r) of pest is <100 descendants per year

Lupinus polyphyllus is a perennial that reproduces sexually but can also reproduce vegetatively via creeping rhizomes below ground (CABI 2018; Ramula 2014). It is pollinated by insects, mainly bees (Jakobsson, Padrón, and Ågren 2015; Pohtio and Teräs 1995). Each plant can have inflorescences with 100-200 flowers that are highly attractive to pollinators, though are nectarless (Pohtio and Teräs 1995; CABI 2018). Hairy pods are produced containing 4-10 seeds that mature in early autumn (CABI 2018; Fremstad 2010).

Lupinus polyphyllus can produce more than 1000 seeds per plant each year, with as many as ~2,500 reported in one study (Aniszewski, Kupari, and Leinonen 2001; OGTR 2013). A European study found an emergence probability of L. polyphyllus seeds from various populations of up to ~40% (Sõber and Ramula 2013). A study in Finland measured establishment of invasive L. polyphyllus populations (i.e., the proportion of seeds that successfully germinated and established) of up to ~35% (Ramula 2014). Therefore, over 800 descendants could result annually.

Extent of Invasion: MODERATE

RANKING Very High >60 countries likely to have established populations of the pest High 30-60 countries likely to have established populations of the pest Moderate 15-29 countries likely to have established populations of the pest Moderately Low 7-14 countries likely to have established populations of the pest Negligible 1-7 countries likely to have established populations of the pest

Lupinus polyphyllus is currently documented in at least seven Minnesota counties (see Proximity to Minnesota).

Portions of all 87 counties would be available based on climate suitability (see Suitability of Minnesota Climate). Suitable habitat estimates suggest county-level distribution would also not be significantly limited by available habitat (see Presence of Hosts). Innate dispersal is unlikely to contribute to long-distance spread of populations (see Innate dispersal capacity).

In two decades in Finland, L. polyphyllus spread an additional 400km (Valtonen, Jantunen, and Saarinen 2006). Extrapolating to Minnesota, a spread of 200km within 10 years from Pine Co, for example, would result in ~35 new counties. Or, if spread were from Becker Co., ~38 new counties could have populations. Therefore, ~45 total counties could have populations (assuming spread from only one county).

6 | L. polyphyllus

Alternatively, future invasion extent could be estimated from the historical county- level reporting in Minnesota. Though the first record of L. polyphyllus in the state was 1970, additional reports did not occur until 1981. From 1981 to the present (2019), 6-8 new counties have reported populations (see Proximity to Minnesota). This amounts to ~0.2 new counties each year (or ~2/decade), or 9-11 total counties in the next 10 years. It is noted that this assumes the current occurrence within the state is adequately surveyed.

Considering the uncertainty from both estimates, an intermediate ranking of ‘moderate’ was selected.

Existence of Vectors: NEGLIGIBLE

RANKING High Vectored by birds or long distance insect migrants Medium Vectored by insects or bats Low Vectored by other mammals Negligible No evidence of any vectors

There is suggestion that L. polyphyllus seeds are dispersed via zoochory by insects and/or rodents (Rask-Jensen 2018), though evidence is lacking.

IMPACT

Problem Elsewhere: MEDIUM

RANKING High Noted as a problem within its native range and areas where it has invaded Medium Noted as a problem only in areas where it has invaded Low Not reported as a problem elsewhere

There is no indication that L. polyphyllus is perceived as a problem in its native western U.S. range.

It is considered a noxious, invasive species in Australia by one report (Randall 2007), though another lists it as a significant weed in natural systems but not as a noxious weed (OGTR 2013). It is generally viewed as problematic throughout Europe and the eastern U.S. (Cao and Sturtevant 2019; CABI 2018; Pergl 2015; Beuthin 2012), though some still regard it as an economically-valuable species within its non-native range (Kurlovich, Stoddard, and Earnshaw 2008). It does not appear to be regulated in any U.S. state (USDA-NRCS 2019).

7 | L. polyphyllus

Impact to Yields and Marketability: LOW

RANKING High >$5 million Medium $5 million to 0.5 million Low <$0.5 million

Lupinus polyphyllus is cultivated for multiple additional purposes, including as a forage and green manure crop (CABI 2018; Cao and Sturtevant 2019). There is one report that hay fields containing L. polyphyllus are more difficult to harvest and have reduced value due to the high alkaloid content of some L. polyphyllus (Fremstad 2010). However, specific estimates of impact could not be found and low-alkaloid cultivars have been developed in effort to mitigate toxicity issues (Kurlovich, Stoddard, and Earnshaw 2008; Aniszewski, Kupari, and Leinonen 2001). Though Minnesota has considered the use of lupin as an alternative forage crop, L. polyphyllus does not appear among the species considered (Putnam 1993). Loss of market value due to L. polyphyllus regulation, or from its impact to other marketable species within Minnesota is therefore unclear and presumed low.

Costs of Quarantine or Other Mitigation (annual): LOW

RANKING High >$5 million Medium $5 million to 0.5 million Low <$0.5 million

Multiple sources state L. polyphyllus can be almost completely removed by properly timed mowing (or grazing) and cuttings removal over multiple years, or with herbicides (e.g., Round-up) (Fremstad 2010; Reinhardt et al. 2003; Valtonen, Jantunen, and Saarinen 2006). However, effective control is also expected to include awareness and regulation of the intentional planting of L. polyphyllus, specifically of mass plantings by transportation authorities or for agriculture or wildlife management (Fremstad 2010; Reinhardt et al. 2003). Complete prohibition is considered unrealistic in Europe (Fremstad 2010).

One estimate from Germany for the cost of L. polyphyllus management via mowing was 300 Euro/hectare annually (Reinhardt et al. 2003). Assuming this is a reasonable estimate for control in Minnesota, under current exchange rates, this is ~$331/ha. Therefore, only 15,000ha (or ~0.07% of state land) would need to be controlled in this manner to nearly $5 million in costs. However, the current extent of infestation does not yet warrant this level of expenditure. We estimate that the economic impact of this weed over the next 10 years for management would be less than $0.5 million per year. As described in the plan, “Elimination of Target Invasive Species - Phase II,” the Minnesota Department of Agriculture received approximately $0.5 million over three years for the management of nine plants designated as “prohibited eradicate”

8 | L. polyphyllus species on the State Noxious Weed List (Chandler 2016). These funds likely underestimate additional expenditures from the MDA and private landowners. Thus, we estimate that $1 million was spent over three years, which amounts to $37,000 per species per year. These calculations assume that action would be taken when naturalized populations are still relatively small.

It is unknown what negative economic impact quarantine of L. polyphyllus may have on the ornamental industry.

Impacts to Recreation or Real Estate (annual): NONE

RANKING High >$5 million Medium $5 million to 0.5 million Low <$0.5 million None $0

Documentation of negative impacts to recreation or real estate by L. polyphyllus could not be found. It is noted that the species is often used for soil stabilization and nutrient improvement (Fremstad 2010), which could positively affect property values and land usage. However, it is unknown how irreplaceable this service is (i.e., if the removal of L. polyphyllus from the market would result in an economic cost to real estate/recreation).

Consequences to Native Species (Score): 3

References are only needed for the “worst case” situation.

RANKING Could reasonably be expected to affect federally listed Threatened and 5 Endangered Species Could directly, negatively impact pollinator 4 Causes local loss of native species 4 Lowers density of native species (empirical support) 3 Infection to native fauna or flora 2 Consumes native fauna or flora 2 Production of toxic substances including allelochemicals 2 Lowers density of native species (presumed due to dense thicket or vining) 2 Host for recognized pathogens/parasites of native species 1 None of the above apply 0

Lupinus polyphyllus is associated with declines in plant species richness across multiple habitat types of its invaded range (Ramula and Pihlaja 2012; Hejda 2013), but effects seem inconsistent. As Ramula and Pihlaja (2012) describe, “However, the lupin was unassociated with differences in plant community composition in terms of species identity or species cover” (Ramula and Pihlaja 2012; p. 2085). This result suggests that while L. polyphyllus may (appear to) exclude a species at a plot scale,

9 | L. polyphyllus the same species is not similarly affected in all plots. Likewise, Hejda (2013) notes: “In the native range, I found no response of native species richness to the cover of Lupinus polyphyllus. In the Czech Republic (central Europe) [where L. polyphyllus is not native], the richness of native species related to it negatively, but the relation was only marginally significant. Contrary to that, the richness of species native to New Zealand [where L. polyphyllus is also not native] related to the cover of Lupinus polyphyllus strongly negatively and the negative relation was significantly stronger than that of species native to Europe.” Available evidence suggests from climates that are most similar to Minnesota, L. polyphyllus may lower the densities of some native species, but will not cause local loss of native species. More observations from climates similar to Minnesota are needed.

Though positive impacts from L. polyphyllus presence has been measured for some pollinators, like bumblebees (Jakobsson, Padrón, and Ågren 2015; Ramula and Sorvari 2017), these effects have only been seen at small spatial scales and do not appear true for all species. For example, invasion of L. polyphyllus decreased the available cover and area of low-growing plant species, which also lowered the plant species richness and diversity in Finland, particularly in verge habitats (Valtonen, Jantunen, and Saarinen 2006). At the same sites, the decreased quality and quantity of larval and adult butterfly host plants resulted in a decrease in the abundance of multiple butterfly and diurnal moth species compared to sites not invaded by L. polyphyllus (Valtonen, Jantunen, and Saarinen 2006). Abundance of other arthropod taxa, including Coleoptera, Diptera, and Hymenoptera, is also lower in sites with L. polyphyllus compared to sites without (Ramula and Sorvari 2017). These effects manifest through changes in the plant community, so they do not constitute a direct effect on pollinators.

The federally-endangered Karner Blue butterfly (Lycaeides melissa samuelsis), a specialist on L. perennis, has been observed ovipositing on L. polyphyllus (Gibbs et al. 2012), but some purport that the larvae cannot develop on it (Summers 2010; Richardson and Jaffe 2018) and L. polyphyllus therefore acts as an ecological sink. Clear empirical evidence could not be found, however, and one study reports that L. melissa samuelsis larvae can develop on L. polyphyllus (Herms et al. 1996).

Leachates from L. polyphyllus litter have been shown to cause delayed and reduced germination of native species in Europe (Loydi et al. 2015). Like other Lupinus spp. (Muzquiz et al. 1994), Lupinus polyphyllus is known to contain and secrete toxic alkaloids in all plant parts, and allelopathic effects from the presence of co- occurrence with L. polyphyllus negatively impacts seedling germination and establishment of other species (Wurst, Vender, and Rillig 2010).

10 | L. polyphyllus

Consequences to Ecosystem Services (Score): 5

RANKING Modification of soil, sediments, nutrient cycling Alteration of genetic resources Alteration of biological control Changes in pollination services Alteration of erosion regimes Affects hydrology or water quality (includes effects of management) Creates a fire hazard Interferes with carbon sequestration

Due to nitrogen-fixing root nodules, L. polyphyllus changes surrounding soil chemistry and can significantly alter nutrient-poor sites through eutrophication (Fremstad 2010; Rask-Jensen 2018).

Lupinus polyphyllus can hybridize with co-occurring Lupinus sp. (see Hybridization/host shift), which changes the local genetic landscape.

In New Zealand, L. polyphyllus was found to alter the plant structuring/succession in river-terrace communities, potentially by accumulating silt material and stabilizing the riparian environment (Holdaway and Sparrow 2006). The clumping growth habits of L. polyphyllus act to trap silt and debris floating within rivers, and has contributed to flooding in some areas as result (Holdaway and Sparrow 2006; Timmins and Mackenzie 1995). Large, clumping deep roots has also resulted in the species being intentionally used for erosion control elsewhere (CABI 2018; Fremstad 2010; Beuthin 2012).

Lupinus polyphyllus produces large amounts of pollen and has tall, showy inflorescences that attract pollinators (Jakobsson, Padrón, and Ågren 2015; Ramula and Sorvari 2017). Though positive impacts from L. polyphyllus presence has been measured for some pollinators, like bumblebees (i.e., Bombus spp) (Jakobsson, Padrón, and Ågren 2015), these effects have only been seen at small spatial scales and do not appear true for all species. For example, invasion of L. polyphyllus decreased the available cover and area of low-growing plant species, which also lowered the plant species richness and diversity in Finland, particularly in verge habitats (Valtonen, Jantunen, and Saarinen 2006). At the same sites, the decreased quality and quantity of larval and adult butterfly host plants resulted in a decrease in the abundance of multiple butterfly and diurnal moth species compared to sites not invaded by L. polyphyllus (Valtonen, Jantunen, and Saarinen 2006). In either case, however, the presence of L. polyphyllus changes the dynamics of local pollination services.

11 | L. polyphyllus

Facilitate Other Invasions: MEDIUM

Invasion by the organism could lead to invasions of other species.

RANKING High The invasive species has facilitated invasions elsewhere The invasive species is a plant or animal that could reasonably be expected Medium to be a host or vector of another invasive species The species has not been reported to facilitate invasion elsewhere and is Low not likely to directly aid in the invasion of other species

A congenur, Lupinus arboreus, has been shown to facilitate the invasion of non-native grasses and forbs to a California coastal prairie community due to its addition of high amounts of nitrogen into the system (Maron and Connors 1996). Similarly, the increased soil nitrogen in old stands of L. nootkatensis has facilitated the establishment of invasive plants in Iceland such as Anthriscus sylvestris (Vetter et al. 2018).

Lupinus polyphyllus is listed as a ”very common” host plant of the invasive moth, Epiphyas postvittana (USDA-APHIS 2009).

REFERENCES

Aniszewski, T., M.H. Kupari, and A.J. Leinonen. 2001. “Seed Number, Seed Size and Seed Diversity in Washington Lupin (Lupinus Polyphyllus Lindl.).” Annals of Botany 87: 77–82. https://doi.org/10.1006/anbo.2000.1300. Beuthin, Mary. 2012. “Plant Guide for Bigleaf Lupine (Lupinus Polyphyllus).” Corvallis, OR: USDA-NRCS Corvallis Plant Materials. https://plants.usda.gov/plantguide/pdf/pg_lupo2.pdf. BRC, Botanical Society of Britain & Ireland and the Biological Records Centre. 2019. “Lupinus Arboreus x Polyphyllus (L. x Regalis).” Online Atlas of the British and Irish Flora. 2019. https://www.brc.ac.uk/plantatlas/plant/lupinus-arboreus-x- polyphyllus-l-x-regalis. Brobäck, David. 2015. “Preventing the Spread of the Invasive Plant Lupinus Polyphyllus.” Uppsala Universitet. http://www.diva- portal.org/smash/get/diva2:893657/FULLTEXT01.pdf. Byleckie, R. 2000. “Wild or Sundial Lupine.” The Rhode Island Wild Plant Society Cultivation Notes 14 (2). http://www.ogtr.gov.au/internet/ogtr/publishing.nsf/Content/biologylupin201 3-toc/$FILE/biologylupin2013-2.pdf. CABI, CAB International. 2018. “Lupinus Polyphyllus (Garden Lupin).” Invasive Species Compendium. Wallingford, UK: CAB International, Invasive Species Compendium. 2018. https://www.cabi.org/isc/datasheet/31710#tosummaryOfInvasiveness. Canadensys. 2019. “Database of Canadian Vascular Plants (VASCAN).” 2019. http://data.canadensys.net/vascan/search. Cao, L., and R. Sturtevant. 2019. “Lupinus Polyphyllus Lindl.” GLANIS (Great Lakes Aquatic Nonindigenous Species Information System). Ann Arbor, MI: NOAA

12 | L. polyphyllus

Great Lake Aquatic Nonindigenous Species Information System, USGS Nonindigenous Aquatic Species Database. 2019. https://nas.er.usgs.gov/queries/greatlakes/FactSheet.aspx?SpeciesID=2715. Chayka, K., and P.M. Dziuk. 2017. “Minnesota Wildflowers.” 2017. https://www.minnesotawildflowers.info/. Clements, J., L. Prilyuk, J. Quealy, and G. Francis. 2008. “Interspecific Crossing among the New World Lupin Species for Lupinus Mutabilis Crop Improvement.” In Lupins for Health and Wealth, Proceedings of the 12th International Lupin Conference, edited by J.A. Palta and J.B. Berger, 324–27. Fremantle, Western Australia: International Lupine Association. EDDMapS, Early Detection and Distribution Mapping System. 2019. “Bigleaf Lupine, Lupinus Polyphyllus Lindl.” Tifton, GA: The University of Georgia - Center for Invasive Species and Ecosystem Health. https://www.eddmaps.org/distribution/uscounty.cfm?sub=12561. Fremstad, Eli. 2010. “Lupinus Polyphyllus.” NOBANIS - Invasive Alien Species Fact Sheet. Online Database of the European Network on Invasive Alien Species. http://www.nobanis.org/files/factsheets/Pacifastacus_leniusculus.pdf. Gibbs, James P., Lawrence B. Smart, Andrew E. Newhouse, and Donald J. Leopold. 2012. “A Molecular and Fitness Evaluation of Commercially Available versus Locally Collected Blue Lupine Lupinus Perennis l. Seeds for Use in Ecosystem Restoration Efforts.” Restoration Ecology 20 (4): 456–61. https://doi.org/10.1111/j.1526-100X.2011.00809.x. Hejda, Martin. 2013. “Do Species Differ in Their Ability to Coexist with the Dominant Alien Lupinus Polyphyllus? A Comparison between Two Distinct Invaded Ranges and a Native Range.” NeoBiota 17: 39–55. https://doi.org/10.3897/neobiota.17.4317. Herms, Catherine Papp, Deborah G. McCullough, Deborah L. Miller, Leah S. Bauer, and Robert A. Haack. 1996. “Laboratory Rearing of Lycaeides Melissa Samuelis (Lepidoptera: Lycaenidae), an Endangered Butterfly in Michigan.” Great Lakes Entomologist 29 (2): 63–75. Holdaway, Robert J., and Ashley D. Sparrow. 2006. “Assembly Rules Operating along a Primary Riverbed-Grassland Successional Sequence.” Journal of Ecology 94 (6): 1092–1102. https://doi.org/10.1111/j.1365-2745.2006.01170.x. Jakobsson, Anna, Benigno Padrón, and Jon Ågren. 2015. “Distance-Dependent Effects of Invasive Lupinus Polyphyllus on Pollination and Reproductive Success of Two Native Herbs.” Basic and Applied Ecology 16 (2): 120–27. https://doi.org/10.1016/j.baae.2014.12.005. Kartesz, J.T. 2015. “The Biota of North America Program (BONAP).” North American Plant Atlas. Chapel Hill, NC: Floristic Synthesis of North America, Version 1.0. http://bonap.net/Napa/TaxonMaps/Genus/County/Lupinus. KEW, Royal Botanic Gardens Kew Science. 2019. “Lupinus Polyphyllus Lindl.” Plants of the World Online. 2019. http://www.plantsoftheworldonline.org/taxon/urn:lsid:ipni.org:names:504939-1. Kurlovich, Boguslav S., Fred L. Stoddard, and Peter Earnshaw. 2008. “Potential and Problems of Lupinus Polyphyllus Lindl.” In ‘Lupins for Health and Wealth’, Proceedings of the 12th International Lupin Conference, edited by J.A. Palta and J.B Berger, 304–7. Fremantle, Western Australia: International Lupine Association. Loydi, A., T. W. Donath, R. L. Eckstein, and A. Otte. 2015. “Non-Native Species Litter Reduces Germination and Growth of Resident Forbs and Grasses: Allelopathic,

13 | L. polyphyllus

Osmotic or Mechanical Effects?” Biological Invasions 17 (2): 581–95. https://doi.org/10.1007/s10530-014-0750-x. Maron, John L., and Peter G. Connors. 1996. “A Native Nitrogen-Fixing Shrub Facilitates Weed Invasion.” Oecologia 105 (3): 302–12. https://doi.org/10.1007/BF00328732. MBG, Missouri Botanical Garden. 2019. “Lupinus x Hybrida.” Plant Finder. 2019. http://www.missouribotanicalgarden.org/PlantFinder/PlantFinderDetails.aspx? kempercode=b764. MN-DNR, Minnesota Department of Natural Resources. 2001. “Riparian Forests in Minnesota: A Report to the Legislature.” http://scholar.google.com/scholar?start=400&q=%22Gap+Analysis+Program%22 +++%22usgs%22&hl=en&as_sdt=0,6#7. ———. 2018. “Minnesota Prairie Conservation Plan: A Habitat Plan for Native Prairie, Grassland and Wetlands in the Prairie Region of Western Minnesota.” Saint Paul, MN: MN DNR. https://doi.org/10.1215/00265667-7137105. ———. 2019. “Why We Need CRP.” Conservation Reserve Program - MN. 2019. https://www.dnr.state.mn.us/crp/index.html. MN-DOT, Minnesota Department of Transportation. 2018. “Railroad Companies Serving Minnesota.” 2018. http://www.dot.state.mn.us/ofrw/railroad/systems.html. MRRA, Minnesota Regional Railroads Association. 2020. “Information about Minnesota Railroads, 2019-2020.” St. Paul, MN. http://www.mnrailroads.com/_pdf/MNRRs-2019-2020.pdf. Muzquiz, Mercedes, Celia De, Carmen Cuadrado, Carmen Burbano, and Rosa Calvo. 1994. “Herbicide-like Effect of Lupinus Alkaloids” 2: 273–80. OGTR, Australian Government Office of the Gene Technology Regulator. 2013. “The Biology of Lupinus L. (Lupin or Lupine).” http://www.ogtr.gov.au/internet/ogtr/publishing.nsf/Content/biologylupin201 3-toc/$FILE/biologylupin2013-2.pdf. Pergl, Jan. 2015. “Lupinus Polyphyllus.” EU Non-Native Organism Risk Assessment Scheme. https://circabc.europa.eu/webdav/CircaBC/env/ias_forum/Library/3 Risk Assessments - check by Scientific Forum/First list update - archive/2 Second submission - for Scientific Forum meeting/Lupinus polyphyllus/Lupinus polyphyllus RA %2B evidence criteria.p. Pohtio, Inari, and Ilkka Teräs. 1995. “Bumblebee Visits to Different Colour Morphs of the Washington Lupine, Lupinus Polyphyllus.” Entomologica Fennica 6 (2–3): 139– 51. https://doi.org/10.33338/ef.83851. Putnam, D.H. 1993. “An Interdisciplinary Approach to the Development of Lupin as an Alternative Crop.” In New Crops, edited by J. Janick and J.E. Simon, 266–77. New York, NY: Wiley. https://hort.purdue.edu/newcrop/proceedings1993/V2- 266.html. Ramula, Satu. 2014. “Linking Vital Rates to Invasiveness of a Perennial Herb.” Oecologia 174 (4): 1255–64. https://doi.org/10.1007/s00442-013-2869-3. Ramula, Satu, and Kati Pihlaja. 2012. “Plant Communities and the Reproductive Success of Native Plants after the Invasion of an Ornamental Herb.” Biological Invasions 14 (10): 2079–90. https://doi.org/10.1007/s10530-012-0215-z. Ramula, Satu, and Jouni Sorvari. 2017. “The Invasive Herb Lupinus Polyphyllus Attracts Bumblebees but Reduces Total Arthropod Abundance.” Arthropod- Plant Interactions 11 (6): 911–18. https://doi.org/10.1007/s11829-017-9547-z. Randall, R.P. 2007. The Introduced Flora of Australia and Its Weed Status. Ecological

14 | L. polyphyllus

Management & Restoration. Vol. 9. Glen Osmond: CRC for Australian Weed Management. Rask-Jensen, Cathrine. 2018. “Seed Dispersal and Phenology of the Invasive Plant Species Bunias Orientalis and Lupinus Polyphyllus in South-East Norway.” Norwegian University of Life Sciences. Reinhardt, F., M.H. Volkswirt, F. Bastiansen, and S. Bruno. 2003. “Economic Impact of the Spread of Alien Species in Germany.” Texte 80-03. Berlin, Germany. http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.135.35&rep=rep1&typ e=pdf. Richardson, M., and D. Jaffe. 2018. “Sundial Lupine, Lupinus Perrenis.” In Native Plants for New England Gardens, 47. Guilford, CT: Globe Pequot. https://books.google.com/books?id=mqVADwAAQBAJ&pg=PA47&lpg=PA47&d q=karner+blue+butterfly+host+plant+%22polyphyllus%22&source=bl&ots=hTbE_5 MKu4&sig=ACfU3U1c4YRz4- JykRK0Wq2hZz9aF7JOqA&hl=en&sa=X&ved=2ahUKEwiT_b2F7OflAhWhSt8KHV 6OAC04ChDoATAIegQICRAB#v=onepag. Sõber, Virve, and Satu Ramula. 2013. “Seed Number and Environmental Conditions Do Not Explain Seed Size Variability for the Invasive Herb Lupinus Polyphyllus.” Plant Ecology 214 (6): 883–92. https://doi.org/10.1007/s11258-013-0216-8. Stace, Clive. 2010. Eu-Dicots, 30. Lupinus L. New Flora of the British Isles. 3rd ed. Cambridge University Press. https://books.google.com/books?id=nJ3YP28EjscC&pg=PA180&lpg=PA180&dq =L.+x+regalis&source=bl&ots=jgJ1uLyDoN&sig=ACfU3U0i9fbrWxtkgWIf1sw3mu mu3gLVBA&hl=en&sa=X&ved=2ahUKEwiq1aqN9OflAhVBPawKHYLbA9YQ6AEw C3oECAYQAQ#v=onepage&q=L. x regalis&f=false. Summers, Carolyn. 2010. “Wildlife in Field, Forest, and Garden.” In Designing Gardens with Flora of the American East, 15. New Brunswick: Rutgers University Press. https://books.google.com/books?id=FMCK_Sl9AWwC&pg=PA15&lpg=PA15&dq =karner+blue+russell#v=onepage&q=karner blue russell&f=false. Timmins, S. M., and I.W. Mackenzie. 1995. “Weeds in New Zealand Protected Natural Areas Database.” Wellington. https://www.doc.govt.nz/globalassets/documents/science-and- technical/docts08.pdf. UMN-Bell, University of Minnesota Bell Museum. 2019. “Minnesota Biodiversity Atlas.” St. Paul, MN: University of Minnesota. https://bellatlas.umn.edu/. USDA-APHIS, (United States Department of Agriculture Animal and Plant Health Inspection Service). 2009. “Light Brown Apple Moth Sterile Insect Field Evaluation Project in Sonoma and Napa, California: Environmental Assessment September 2009.” Riverdale, MD. USDA-ARS, United States Department of Agriculture - Agricultural Research Service. 2012. “USDA Plant Hardiness Zone Map.” 2012. https://planthardiness.ars.usda.gov/PHZMWeb/. USDA-NRCS, United States Department of Agriculture Natural Resources Conservation Service. 2019. “Plant Profile: Lupinus Polyphyllus Lindl., Bigleaf Lupine.” Plants Database. 2019. https://plants.usda.gov/core/profile?symbol=LUPO2. Valtonen, Anu, Juha Jantunen, and Kimmo Saarinen. 2006. “Flora and Lepidoptera Fauna Adversely Affected by Invasive Lupinus Polyphyllus along Road Verges.” Biological Conservation 133 (3): 389–96. https://doi.org/10.1016/j.biocon.2006.06.015.

15 | L. polyphyllus

Vetter, Vanessa M.S., Nils B. Tjaden, Anja Jaeschke, Constanze Buhk, Veronika Wahl, Pawel Wasowicz, and Anke Jentsch. 2018. “Invasion of a Legume Ecosystem Engineer in a Cold Biome Alters Plant Biodiversity.” Frontiers in Plant Science 9 (June): 1–12. https://doi.org/10.3389/fpls.2018.00715. WI-DNR, Wisconsin Department of Natural Resources. 2019. “Big-Leaf Lupine (Lupinus Polyphyllus).” Invasive Species. 2019. https://dnr.wi.gov/topic/Invasives/fact/BigleafLupine.html. Williams, J. 2005. “Minnesota Highway Mileage.” St. Paul, MN: Research Department of the Minnesota House of Representatives. https://www.house.leg.state.mn.us/hrd/pubs/ss/ssmnhm.pdf. Wohlman, Matthew. 2017. “Minnesota Department of Agriculture, House Committee on Agriculture Finance.” St. Paul, MN. https://www.senate.mn/committees/2017- 2018/3088_Committee_on_Agriculture_Rural_Development_and_Housing_Finan ce/2017 MDA Budget Presentation 1-30-17 mh.pdf. Wurst, Susanne, Veruschka Vender, and Matthias C. Rillig. 2010. “Testing for Allelopathic Effects in Plant Competition: Does Activated Carbon Disrupt Plant Symbioses?” Plant Ecology 211 (1): 19–26. https://doi.org/10.1007/s11258-010- 9767-0.

16 | L. polyphyllus